Spotted hyena

Name and taxonomy

Johann Erxleben gave the species its scientific name in 1777. He worked from a description that Thomas Pennant published in 1771, based on a captive animal. Pennant was the first writer to separate the spotted hyena from the striped hyena in a consistent way. Crocuta became a genus of its own in 1828. The name Crocuta traces back through Greek to Sanskrit words for the golden jackal, not to a Latin word for saffron.

The spotted hyena sits in the family Hyaenidae within the order Carnivora. Its closest relatives are the striped hyena, the brown hyena and the aardwolf. Naturalists once named many subspecies because coat and size vary so much across Africa. In 1939 L. Harrison Matthews showed that skulls from one Tanzanian population held all of that variation. Today the species is treated as a single form with no subspecies.

Evolution and relatives

The line that led to Crocuta split from the striped and brown hyenas about 10 million years ago. The oldest fossils of the genus come from Africa, about 3.63 to 3.85 million years ago. The earliest remains of the living species are from Sterkfontein in South Africa, at least 2.1 million years old. Fossil clans probably began to form about 350,000 years ago, when other predators competed hard for carcasses. Sharp cutting teeth behind the bone-crushing premolars turned the species from a scavenger into a pack hunter.

Crocuta spread out of Africa and lived across Europe and Asia for at least a million years. These cave hyenas are usually treated as a subspecies of the spotted hyena. Their nuclear genome suggests the African and Eurasian groups split about 2.5 million years ago and still interbred afterwards. Cave hyenas vanished from Western Europe between 14 and 11,000 years ago, as grassland gave way to woodland. Paintings in the Lascaux and Chauvet caves show that they kept their spots.

Appearance

The spotted hyena is built heavy at the front, with a thick neck, a wide flat head and a blunt muzzle. The forelegs outreach the hind legs, so the back slopes down to a rounded rump. The ears are rounded, unlike the pointed ears of the other hyenas. Each foot has four toes with short blunt claws that do not retract. Adults measure 95 to 150 centimetres from head to rump and stand 70 to 91.5 cm at the shoulder. The short tail measures 30 to 36 cm and carries a black brush of hair.

Adults usually weigh 45 to 80 kilograms, and the heaviest on record reached 90 kg. The female is the larger sex, which is rare among mammals. Males weigh about 45 to 60 kg, and females weigh 55 to more than 70 kg. The heart makes up close to 1% of body weight, which supports long chases. The skull is among the strongest in the Carnivora for its size, and the bite reaches 4,500 newtons in one measured animal.

The coat is sandy, yellowish or grey with an irregular pattern of dark spots on the back, flanks and legs. The spots are reddish, deep brown or nearly black and are commonly about 20 millimetres across. They are darkest in young animals and fade with age. Researchers use the spot pattern to identify individuals. The fur is short and coarse, with no long mane along the spine. The female's clitoris is shaped like a penis and can become erect, and fused labia form a false scrotum.

Habitat and range

The spotted hyena lives in savanna, semi-desert, open and dry woodland, swamps and mountain forest. It is recorded up to 4,000 metres above sea level in East Africa and Ethiopia. It stays out of lowland rainforest, the highest alpine zones and the driest desert. In West Africa it keeps to the Guinea and Sudan savannas and avoids the coastal forest belt.

The species once lived across almost all of sub-Saharan Africa. Its range is now patchy, most of all in West Africa, and populations gather in and around protected areas. Large continuous populations remain in Ethiopia, Kenya, Tanzania, Botswana, Namibia and the lowveld of South Africa. It has been wiped out in Algeria and Lesotho and from much of southern South Africa. GBIF holds records from 71 countries, but that count includes zoo animals and does not describe the native range.

Density varies from 0.006 to 1.7 animals per square kilometre. The richest plains of Kenya and Tanzania hold more than one hyena per square kilometre. In good habitat the spotted hyena outnumbers every other large carnivore. In desert the striped and brown hyenas are more common than it is.

Diet and hunting

The spotted hyena is the most carnivorous hyena and mainly a predator. Studies since the 1960s show that it kills 60% to 95% of its own food, with the rest scavenged. It prefers prey of 56 to 182 kilograms and takes whatever is common where it lives. In East Africa that means wildebeest, zebra, gazelles and topi. In Kruger the main prey is impala, and in the dry south-west it is gemsbok and springbok. It also eats termites, crocodile eggs, ostrich eggs, hares and the young of giraffe, hippo and rhino.

Hyenas often run through a herd to pick out a young, old or weak animal. They then chase it for several kilometres at up to 60 kilometres per hour. One chase in the Kalahari covered 24 km before an eland was caught. Most hunting parties hold one or two animals, but zebra hunts draw larger groups. A single hyena can pull down a wildebeest three times its own weight. There is no killing bite: the hunters pull the prey down and eat it as it dies.

Feeding is a race, because the noise draws the rest of the clan. A group of 20 to 30 hyenas can strip an adult wildebeest to a pile of stomach contents in 13 minutes. One animal can swallow food equal to 25% to 30% of its body weight at a sitting. The gut digests skin, bone and hooves, so only the horns and stomach contents are left. In the Serengeti a nursing female may commute 80 km round trip to reach migrating herds.

Behaviour and social life

Clans hold from 3 to 80 members, and clan size follows the local prey supply. A clan shares a territory and a communal den, but its members forage alone or in small groups most of the time. The whole clan gathers only at a kill, at the den or when it defends its border. Females stay in their birth clan for life, and males leave at about two and a half years. Immigrant males join a queue and move up the ranks only as older males die.

Every adult female outranks every immigrant male, and a low-ranking female can push a high-ranking male off a kill. A cub takes its mother's rank, one step below her, and keeps it as an adult. Rank depends on allies, not on size or strength, much as it does among baboons. High-ranking females eat first, breed earlier and raise more surviving cubs. A hyena that falls below its inherited rank in its first months is about 1.5 times more likely to die.

Territory size runs from under 40 square kilometres in the Ngorongoro Crater to over 1,000 in the Kalahari. Clans mark the border with anal-gland paste on grass stems and with shared latrines. A hunting hyena may stop dead when its prey crosses into the next clan's land. The species is mostly active at night and around dusk and dawn. Hyenas in Kenya were active for about 32% of the day and walked more than 12 km in 24 hours.

Communication and senses

The spotted hyena has one of the widest vocal ranges of any carnivore. The long rising whoop is a contact call and can be heard far across the plain. The giggle, the famous laugh, comes from a hyena that is afraid or excited, often while it is being chased. A low mooing call comes from an impatient animal waiting at a kill. Grunts, growls and yells go with aggression. A 2022 study found that each hyena's whoop carries an individual signature.

Clan-mates greet by standing head to tail and sniffing each other's raised genitals for up to 30 seconds. An erection during the greeting is a sign of submission, not dominance. A submissive hyena also flattens its ears, lowers its rump and may walk on its front knees. The anal glands produce a paste that smells of burning or cheap soap, and people can smell it several metres downwind. Hyenas can tell clan members from strangers by this scent, which is shaped by the bacteria in the gland. Like other hunters of the night, the species relies on keen smell, hearing and sight to find prey and carrion.

Reproduction and life cycle

Spotted hyenas breed in any month, with a birth peak in the wet season. Both sexes mate with several partners, and no pair bond forms. A male approaches a female in heat with bows and paws the ground, and he retreats at once if she turns on him. Females favour younger males that joined the clan after they themselves were born. Mating is awkward, because the male must enter through the female's clitoris, so forced mating is impossible.

Gestation lasts about 110 days. The female gives birth through her clitoris, which tears and takes weeks to heal. The usual litter is two cubs, and the range is 1 to 4. A newborn weighs about 1 to 1.5 kilograms and is born black, with open eyes and erupted teeth. Cubs of the same sex often fight at once, and the weaker one may die. An estimated 25% of hyenas die this way within a month of birth.

The mother moves her cubs to the communal den at 2 to 5 weeks. She alone feeds them, on milk richer in protein and fat than that of any other land carnivore. Weaning comes late, at 12 to 16 months, when the cubs already have their adult teeth. Cubs join hunts after their first year, and both sexes are mature at about three years. Wild animals live up to 20 years, and one captive hyena reached 41. The average zoo lifespan is 12 years.

Predators and threats

Lions are the main natural killer of spotted hyenas. In one study 13 of 24 hyena carcasses had been killed by lions, and in Etosha lions cause up to 71% of hyena deaths. A male lion may maul a hyena and leave it uneaten. Hyenas answer by mobbing lions that enter their territory. African rock pythons have been seen to take adult hyenas, and hyenas keep clear of crocodile water.

Half of all cubs die before puberty, most of them just after weaning. Hyenas from a neighbouring clan, and sometimes females of the same clan, kill cubs. The species carries antibodies to rabies, canine distemper and several other diseases. A distemper outbreak in 1993 to 1994 struck both hyenas and lions in the Serengeti. Outside parks people are the biggest killer: ranchers shoot, snare, spear and poison hyenas, and cars kill them on roads.

Population and conservation

The IUCN Red List rates the spotted hyena as Least Concern. Its wild population is estimated at 27,000 to 47,000, with several subpopulations above 1,000. The largest numbers live in Kenya, Tanzania, South Africa and Namibia.

Numbers are falling fast outside protected areas because of persecution and habitat loss. The species is listed as threatened in Benin, Burundi, Cameroon, Mauritania, Niger, Nigeria, Rwanda and Sierra Leone. Its survival depends more and more on parks and reserves. The IUCN hyena specialist group names the animal's bad public image as a threat in itself. Many people think it is not worth saving, so education is part of the conservation plan.

Relationship with humans

The spotted hyena is shy of people by day and flees from up to 300 metres away. At night it grows bolder and may follow a person closely. Attacks on people are rare but probably under-reported. Victims are most often women, children and sick men who sleep outside in the hot season. Two large hyenas killed 27 people around Mlanje in Malawi in 1962. A 2004 report counted 35 deaths in one year along a 20 km stretch of road in Mozambique.

Hyenas eat human corpses readily. Maasai and Hadza tradition leaves the dead in the open for hyenas to take. A body the hyenas refuse brings disgrace, so families may smear it with ox fat and blood. Addis Ababa holds up to a thousand hyenas that live on rubbish and stray dogs. In 2013 hyenas killed a baby in the city, and a cull that year shot ten of them.

The spotted hyena takes cattle, sheep and goats, and in parts of Tigray it prefers donkeys. In northern Kenya 90% of stock losses happened outside thorn fences. Hunters seldom want it as a trophy, and most African peoples refuse to eat its meat. Body parts, especially the tail, paws and genitals, are sold for traditional medicine in several countries. It is easy to tame but almost impossible to house-train, and one tame animal tore up a nailed plank in the Tower of London.

In culture and myth

Across Africa the spotted hyena is seen as sly, greedy, foolish and dangerous, but also as powerful and sacred. In many West African tales it stands for dirty habits and the reversal of normal life. Among the Beng of Côte d'Ivoire it plays a bad Muslim who mocks the local spirits. The Beng also leave a village if a hyena defecates inside it. The Tabwa of East Africa say a hyena carried the sun down to warm a cold world.

In the Gelede cult of the Yoruba, a hyena mask closes the èfè ceremony at dawn, because the hyena ends every meal. The Korè cult of the Bambara in Mali treats the hyena as a being of both sexes that joins male and female in the initiate. In Harar, Ethiopia, townspeople feed hyenas by hand and believe they keep devils away. Across East and West Africa, people say that witches ride hyenas after dark and that sorcerers take hyena form. The Ethiopian name for such a were-hyena is bouda.

Western ideas start with Aristotle, who called the animal cowardly and a corpse eater but rejected the hermaphrodite story. Pliny the Elder added that it could copy human voices and that its body parts cured disease. The medieval Physiologus made it a grave-robber with both sexes. In the 20th century Hemingway's Green Hills of Africa and Disney's The Lion King added gluttony and stupidity. Hyena biologists protested the film, and one researcher sued the studio.